Although the mere fact that parthenogenesis occurs at all is, in my opinion, sufficient to disprove the theory of rejuvenescence, it is well to remember that parthenogenesis is now the only method of reproduction in many species (although we do not know the period of time over which these conditions have extended), and is nevertheless unattended by any perceptible decrease in fertility.
From all these considerations we may draw the conclusion that the process of rejuvenescence, as described above, cannot be accepted either as the existing or the original meaning of conjugation, and the question naturally arises as to what other significance this latter process can have possessed at its first beginning.
Rolph[[187]] has expressed the opinion that conjugation is a form of nutrition, so that the two conjugating individuals, as it were, devour each other. Cienkowsky[[188]] also regards conjugation as merely ‘accelerated’ assimilation. There is, however, not only an essential difference but a direct contrast between the processes of conjugation and nutrition. With regard to Cienkowsky’s view, Hensen[[189]] has well said that ‘coalescence in itself cannot be an accelerated nutrition, because even if we admit that both individuals are in want of nourishment, it is impossible that the need can be supplied by this process, unless one of them perishes and is really devoured.’ In order that an animal may serve as the food of another, it must perish and must be brought into a fluid form, and finally it must be assimilated. In the case before us, however, two protoplasmic bodies are placed side by side and coalesce, without either of them passing into the liquid state. Two idioplasms unite, together with all the hereditary tendencies contained in them; but although it is certain that nutrition in the proper sense of the word cannot take place, because neither of the animals receives an addition of liquid food by the coalescence, yet the consequence of this process must be in one respect similar to that of nutrition and growth:—the mass of the body and the quantity of the forces contained in it undergo simultaneous increase. It is not inconceivable that effects are by this means rendered possible, which under the peculiar circumstances leading to conjugation, could not have been otherwise produced.
I believe that this is at any rate the direction in which we shall have to seek for the first meaning of conjugation and for its phyletic origin. This first result and meaning of conjugation may be provisionally expressed in the following formula:—conjugation originally signified a strengthening of the organism in relation to reproduction, which happened when from some external cause, such as want of oxygen, warmth, or food, the growth of the individual to the extent necessary for reproduction could not take place.
This explanation must not be regarded as equivalent to that afforded by the theory of rejuvenescence; for the latter process is said to be necessary for the continuance of reproduction, and ought therefore to occur periodically quite independently of external circumstances; while according to my theory, conjugation at first only occurred under unfavourable conditions, and assisted the species to overcome such difficulties.
But whatever the original meaning of conjugation may have been, it seems to have become already subordinated in the higher Protozoa, as is indicated by the changes in the course taken by this process. The higher Protozoa when conjugating do not as a rule coalesce completely and permanently[[190]] in the manner followed by the lower Protozoa, and it seems to me possible, or even probable, that in the former the process has already gained the full significance of sexual reproduction, and is to be looked upon as a source of variability.
Whether this be so or not, I believe it is certain that sexual reproduction could not have been entirely abandoned at any period since the time when the Metazoa and Metaphyta first arose; for they derived this form of reproduction from their unicellular ancestors.
We know that organs and characters which have persisted through a long series of generations are transmitted with extreme tenacity, even when they have ceased to be of any direct use to their immediate possessors. The rudimentary organs in various animals, and not least in man, afford very strong proofs of the soundness of this conclusion. Another example has only recently been discovered in the sixth finger, which has been shown to exist in the human embryo[[191]], a part which has only been present in a rudimentary form ever since the origin of the Amphibia[[192]]. Superfluous organs become rudimentary very slowly, and enormous periods must elapse before they completely disappear, while the older a character is, the more firmly it becomes rooted in the organism. What I have above called the physical constitution of a species is based upon these facts, and upon them depend the tout ensemble of inherited characters, which are adapted to one another and woven together into a harmonious whole. It is this specific nature of an organism which causes it to respond to external influences in a manner different from that followed by any other organism, which prevents it from changing in any way except along certain definite lines of variation, although these may be very numerous. Furthermore these facts ensure that characters cannot be taken at random from the constitution of a species and others substituted for them. Such a variation as a mammal wanting the firm axis of the backbone is an impossibility, not only because the backbone is necessary as a support to the body, but chiefly because this structure has been inherited from times immemorial, and has become so impressed upon the mammalian organization that any variation so great as to threaten its very existence cannot now take place. The view here set forth of the origin of hereditary variability by amphigonic reproduction, makes it clear that an organism is in a state of continual oscillation only upon the surface, so to speak, while the fundamental parts of its constitution, which have been inherited from extremely remote periods, remain unaffected.
Thus sexual reproduction itself did not cease after it had existed in the form of conjugation through innumerable generations of the vast numbers of species which have been included under the Protozoa; it did not cease even when its original physiological significance had lost its importance, either completely or in part. This process, however, had come to possess a new significance which ensured its continuance, in the enormous advantage conferred on a species by the power of adapting itself to new conditions of life, a power which could only be preserved by means of this method of reproduction. The formation of new species which among the lower Protozoa could be achieved without amphigony, could only be attained by means of this process in the Metazoa and Metaphyta. It was only in this way that hereditary individual differences could arise and persist. It was impossible for amphigony to disappear, for each species in which it was preserved was necessarily superior to those which had lost it, and must have replaced them in the course of time; for the former alone could adapt itself to the ever-changing conditions of life, and the longer sexual reproduction endured, the more firmly was it necessarily impressed upon the constitution of the species, and the more difficult its disappearance became.
Sexual reproduction has nevertheless been lost in some cases, although only at first in certain generations. Thus in the Aphidae and in many lower Crustacea, generations with parthenogenetic reproduction alternate with others which reproduce themselves by the sexual method. But in most cases it is clear that this partial loss of amphigony conferred considerable advantages upon the species by giving increased capabilities for the maintenance of existence. By means of partial parthenogenesis a much more rapid increase in the number of individuals could be attained in a given time, and this fact is of the highest importance for the peculiar circumstances under which these species exist. A species of Crustacean which inhabits rapidly drying pools, and developes from winter-eggs which have remained dried up in the mud, has, as a rule, only a very short time in which to secure the existence of succeeding generations. The few sexual eggs which have escaped the attacks of numerous enemies develope immediately after the first shower of rain; the animals attain their full size in a few days and reproduce themselves as virgin females. Their descendants are propagated in the same manner, and thus in a short time almost incredible numbers of individuals are formed, until finally the sexual eggs are again produced. If now the pool dries up again, the existence of the colony is secured, for the number of animals which produce sexual eggs is very large, and the eggs themselves are of course far more numerous, so that in spite of the destructive agencies to which they are subjected, there will be every chance of the survival of a sufficient number to produce a new generation at a later period. Here, therefore, sexual reproduction has not been abandoned accidentally or from any internal cause, but as an adaptation to certain definite necessities imposed upon the organism by its surroundings.